Introduction

Glyptophidium effulgens was originally de-scribed by Nielsen and Machida (1988) on thebasis of 3 specimens from off the Philippines.Nielsen and Machida (1988) distinguished itfrom other congeners by the following combina-tion of characters: pelvic-fin ray 1, precaudal ver-tebrae 11, median branchial tooth patch 1, orbitdiameter 29.0–31.0% of head length, pseudo-branchial filaments 7–8, and a relatively thinsagitta with a distinct depression in the dorsalrim. Since their original description, additionalspecimens of the species have not been collected.

We have recently had an opportunity to exam-ine the type specimens of Glyptophidium efful-gens and types of G. lucidum in detail, leading usto a conclusion that G. effulgens is a junior syn-onym of G. argenteum Alcock, 1889.

Materials and Methods

Counts and measurements follow Hubbs andLagler (1958) and Nakae and Matsuura (2009).The tail of Glyptophidium is fragile and easily

damaged during collecting operations. As a re-sult, many museum specimens lack the caudaltip. In the following pages, the standard length ofspecimens lacking the caudal tip is shown with“�” after the value presented. Sagittas were ob-served by computerized tomography (CT) scan(ALOKA LCT-100). Institutional abbreviationsfollow Leviton et al. (1985). Specimens exam-ined in this study are listed below.

Glyptophidium argenteum (20 specimens):USNM 99158 [201� mm standard length (SL),holotype of G. effulgens], Albatross sta. D-5410(10°28�45�N, 124°05�30�E), 704 m, bottomtrawl, 17 Mar. 1909; USNM 272001 (210� mmSL, paratype of G. effulgens), same data asUSNM 99158; ZMUC P77783 (228� mm SL,paratype of G. effulgens), same data as USNM99158; USNM 99057 (205� to 244 mm SL, 3specimens), Albatross sta. D-5410 (10°28�45�N,124°05�30�E), 704 m, Agassiz trawl, 18 Mar.1909; USNM 99160 (179� to 217� mm SL, 3),Albatross sta. 5405 (10°49�20�N, 124°24�23�E),479 m, Agassiz trawl, 17 Mar. 1909; USNM99179 (171 to 229� mm SL, 3), Albatross sta.D-5373 (13°40�N, 121°31�10�E), 618 m, beam

Glyptophidium effulgens Nielsen and Machida, 1988, a junior synonym of Glyptophidium argenteum Alcock, 1889

(Actinopterygii, Teleostei, Ophidiiformes)

Masanori Nakae* and Keiichi Matsuura**

Collection Center, National Museum of Nature and Science, 3–23–1 Hyakunin-cho, Shinjuku-ku, Tokyo, 169–0073 Japan

*E-mail: nakae@kahaku.go.jp**E-mail: matsuura@kahaku.go.jp

(Received 30 November 2009; accepted 18 December 2009)

Abstract The type specimens of Glyptophidium effulgens Nielsen and Machida, 1988 were com-pared with specimens of G. argenteum Alcock, 1889 and G. lucidum Smith and Radcliffe in Rad-cliffe, 1913, including types of the latter. These comparisons revealed that G. effulgens is a juniorsynonym of G. argenteum. One of the paratypes of G. lucidum is also identifiable as G. argenteum.Key words : Actinopterygii, Ophidiiformes, Ophidiidae, synonymy, Glyptophidium effulgens,Glyptophidium argenteum, Glyptophidium lucidum.

Bull. Natl. Mus. Nat. Sci., Ser. A, 36(1), pp. 27–32, February 22, 2010

trawl, 2 Mar. 1909; ZMUC P77772–P77777(134� to 234� mm SL, 6), Galathea sta. 436(10°12�N, 124°14�E), 710 m, sledge trawl, 9Aug. 1951; NSMT-P 78388 (93.4� mm SL), offHeda, Suruga Bay, Shizuoka Prefecture, Honshu,Japan, 300–400 m, bottom trawl, 1 Apr. 1986;USNM 99913 [194 mm SL, paratype of G. lu-cidum (see “Results and Discussion” below)], Al-batross sta. 5624 (0°12�15�N, 127°29�30�E),527 m, trawl, 29 Nov. 1909.

Glyptophidium lucidum (21): USNM 74144(211� mm SL, holotype), Albatross sta. 5625(0°07�N, 127°28�E), 421 m, beam trawl, 29 Nov.1909; USNM 99107 (149� to 182� mm SL, 7,paratypes), Albatross sta. D-5626 (0°07�30�N,127°29�E), 485 m, trawl, 29 Nov. 1909; ZMUCP77548 (157� mm SL, paratype), Albatross sta.D-5626 (0°07�30�N, 127°29�E), 485 m, trawl, 29Nov. 1909; USNM 99272 (91.1� to 195 mm SL,11), Albatross sta. 5625 (0°07�N, 127°28�E),421 m, beam trawl, 29 Nov. 1909; NSMT-P48338 (149� mm SL), South Senoumi Bank (34°45�37�N, 138°29�32�E–34°45�58�N,138°28�23�E), Suruga Bay, Shizuoka Prefecture,Honshu, Japan, 240–410 m, bottom trawl, 4 Oct.1995.

Results and Discussion

In their review of Glyptophidium, Nielsen andMachida (1988) recognized 7 species includingtheir new species G. effulgens from the Philip-pines. They stated that G. effulgens (Fig 1A) isdistinguishable from the closely related species,G. argenteum (Fig. 1B) and G. lucidum in havingthe following characters: orbit diameter 29.0–31.0% of head length, pseudobranchial filaments7–8, and a relatively thin sagitta with a distinctdepression in the dorsal rim. However, our exam-ination of the type specimens of G. effulgens re-vealed that it is indistinguishable from G. argen-teum as values for almost all meristic and mor-phometric characters (including orbit diameter)overlap (Fig. 2, Table 1); both have gill rakers onthe anterior gill arch that are long and thin withseveral knobs on the inner surface (Fig. 3); theshape of the dorsal rim of the sagitta and itsthickness (26.5–28.0% of sagitta width in G. ef-fulgens vs. 26.5–36.2% in G. argenteum) arevariable and thus not useful for separating thetwo species (Fig. 4; see also figures of Nakae andMatsuura, 2009). In addition, the distribution ofG. effulgens completely overlaps that of G. ar-

28 Masanori Nakae and Keiichi Matsuura

Fig. 1. Holotype of Glyptophidium effulgens, USNM 99158, 201 mm�SL (A) and specimen of G. argenteum,USNM 99057-3407, 208 mm SL (B).

Synonymy of Ophidiids 29

Fig. 2. Comparisons of orbit diameter versus body depth at anal-fin origin in specimens of Glyptophidium ar-genteum, G. effulgens, G. lucidum and a paratype of G. lucidum USNM 99113. Three damaged specimensexamined (1 argenteum, 2 lucidum) are excluded. Abbreviations: BDA, body depth at anal-fin origin; HL,head length; OD, orbit diameter; P1, pectoral-fin ray; PSB, pseudobranchial filaments.

Table 1. Meristic and morphometric characters of 3 species of Glyptophidium (argenteum species-group) fishes. Datafrom Nielsen and Machida (1988) in parentheses.

Species G. argenteum G. effulgens G. lucidum

Specimens 17 specimens from USNM 99158 USNM 272001 ZMUC P77783 21 specimens from USNM and ZMUC Holotype Paratype Paratype USNM and ZMUC

Standard length (mm) 93.4*–244* (127–287) 201* (202) 210* (212*) 228 (228) 91.1*–211* (115–196)Dorsal-fin rays 143–155 (140–158) 144* (141) 107* (107*) 142 (141) 133–140 (133–146)Anal-fin rays 118–130 (114–133) 122* (124) 89* (89*) 117 (125) 110–122 (109–122)Pectoral-fin rays 20–24 (20–24) 21 (21) 22 (22) 22 (22) 23–26 (23–26)Pelvic-fin rays 1 (1) 1 (1) 1 (1) 1 (1) 1 (1)Caudal-fin rays 6–7 (6–7) – (–) – (–) – (–) 7–8 (7–8)Gill rakers on anterior arch: 1–4�17–21�4–7 1�18�6 2�18�6 3�19�6 1–4�17–20�5–7

dorsal small�long�ventral small (0–5�17–23�3–8) (1�19�5) (2�19�6) (3�19�6) (0–3�14–20�4–9)Total gill rakers on anterior arch 24–28 (23–31) 25 (25) 26 (27) 28 (28) 24–28 (23–27)Median basibranchial tooth patch 1 (1) 1 (1) 1 (1) 1 (1) 1 (1)Precaudal vertebrae 11 (11–12) 11 (11) 11 (11) 11 (11) 11 (11)Caudal vertebrae 78–82 (74–85) 74 (74) 51* (51*) 74 (73) 72-73 (70–81)Dorsal-fin origin above vertebra number 3–4 (2–4) 3 (3) 3 (3) 3 (3) 3–4 (3–4)Anal-fin origin below dorsal ray number 19–21 (18–23) 21 (20) 19 (20) 19 (21) 19–23 (20–24)Anal-fin origin below vertebra number 13–15 (13–16) 14 (14) 13 (13) 13 (14) 13–15 (12–15)Pseudobranchial filaments 7–12 (7–12) 7 (7) 9 (8) 9 (8) 12–15 (11–15)

In % of head lengthOrbit diameter 27.1–30.6 (21.5–28.5) 28.5 (29.5) 28.5 (29) 29.3 (31) 31.4–34.6 (31.5–40.5)Body depth at dorsal-fin origin 66.4–85.4 (66–91) 76.3 (80) 76.8 (78) 76.2 (81) 71.9–81.3 (75–88)Body depth at anal-fin origin 56.4–67.1 (52–69) 62.8 (63) 65.4 (64) 63.2 (65) 57.5–70.3 (61–77)

In % of preanal lengthHead length 60–68.3 (55–74) 63.6 (–) 65.9 (–) 64.8 (–) 56.7–66.1 (53–66)

Condition of gill rakers long and thin long and thin long and thin long and thin short and robust

*Caudal tip lacking.

genteum (see Nielsen and Machida, 1988).Although, Nielsen and Machida (1988) stated

that the holotype of G. argenteum Alcock, 1899

is in a very poor condition and only 19 of 118specimens examined by them have a completetail, their description showed no doubts about its

30 Masanori Nakae and Keiichi Matsuura

Fig. 3. Gill raker on angle between epi- and ceratobranchial on left first gill arch in three nominal Glyptophidi-um species. A, G. argenteum, ZMUC-P 77772, 228 mm SL; B, holotype of G. effulgens, USNM 99158,201� mm SL; C, G. lucidum, USNM 99109, 212� mm SL. [Figures of G. argenteum and lucidum slightlymodified from Nielsen and Machida (1988).]

Fig. 4. Medial view of sagitta of Glyptophidium argenteum (A), effulgens (B), and lucidum (C). Bar 2 mm.[Figures modified from Nielsen and Machida (1988).]

taxonomic status.Nielsen and Machida (1988) stated that the

type specimens of G. effulgens are good in condi-tion except for the broken pelvic-fin rays and themissing caudal tip of 1 paratype (USNM272001). However our examination of the typespecimens of G. effulgens revealed that the holo-type (USNM 99158) also lacks the caudal tip.We were able to determine from X-ray pho-tographs of the holotype (Fig. 5) that the posteri-or ends of the dorsal and anal-fins are missing,and making it impossible accurately to count dor-sal and anal-fin rays. Several of Nielsen andMachida’s (1988) fin ray and other meristiccounts for the holotype and intact paratype(ZMUC P77783) differ from ours: dorsal-fin raysof Nielsen and Machida 141 in both the holotypeand paratype vs. 142-144, anal-fin rays of

Nielsen and Machida 124–125 vs. 117–122, cau-dal vertebrae of Nielsen and Machida 73–74 vs.74 in both the holotype and paratype, total gillrakers of Nielsen and Machida 25–28 vs. 25–27and pseudobranchial filaments of Nielsen andMachida 7–8 vs. 7–9 by us. As our values for allof these characters were based on X-rays, we areconfident in their accuracy.

One paratypes of G. lucidum, USNM 99113,has 24 pectoral-fin rays, 8 pseudobranchial fila-ments, relatively long and thin rakers on the ante-rior gill arch, and orbit diameter 28.9% of headlength (Fig. 2) clearly identifying it as G. argen-teum.

Nielsen and Machida (1988) counted 24 gillrakers on the anterior arch in the holotype of G.lucidum while Smith and Radcliffe (1913) stated25 in their original description. Our examination

Synonymy of Ophidiids 31

Fig. 5. X-ray photographs of caudal tip of the type specimens of Glyptophidium effulgens. A, USNM 99158(holotype); B, USNM 272001 (paratype); C, ZMUC P77783 (paratype).

of the holotype supports Smith and Radcliffe(1913). On the other hand, Smith and Radcliffe(1913) counted about 10 pseudobranchial fila-ments in the holotype, but Nielsen and Machida(1988) found 15, which we confirmed.

Acknowledgments

We express our sincere thanks to Jeffrey T.Williams (USNM), Jørgen G. Nielsen andTammes Menne (ZMUC) for the loan of speci-mens, Sandra J. Raredon (USNM) for taking dig-ital X-ray images of the type specimens of G. ef-fulgens, and Kyoko Iwami (NSMT) for her CTscan operation. We thank Martin F. Gomon (Mu-seum Victoria) for critically reading the manu-script and offered helpful comments, and Yoshi-hiko Machida (Kochi University) and Gento Shi-nohara (NSMT) for their useful comments. Thisstudy was partially supported by Grant in Aidsfor Scientific Research (A) by the Japan Societyfor the Promotion of Science, Japan (19208019).

References

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Hubbs, C. L. and K. F. Lagler 1958. Fishes of the GreatLakes Region. Bulletin of the Cranbrook Institute ofScience, 26: i–xi�1–186.

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32 Masanori Nakae and Keiichi Matsuura